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UNIVERSITI PUTRA MALAYSIA
POPULATION ECOLOGY, REPRODUCTIVE BEHAVIOUR AND FEEDING HABIT OF Helopeltis antonii Signoret ON CASHEW
(Anacardium occidentale L.) PLANTS
SISWANTO
FP 2007 28
POPULATION ECOLOGY, REPRODUCTIVE BEHAVIOUR AND FEEDING HABIT OF Helopeltis antonii Signoret ON CASHEW (Anacardium
occidentale L.) PLANTS
By
SISWANTO
Thesis Submitted to the School of Graduate Studies, Universiti Putra Malaysia, in Fulfilment of the Requirements for the Degree of
Doctor of Philosophy
November 2007
DEDICATION
I dedicate this thesis to my wife Yuyun Nurohmah and children Arina
Yusianti, Naufal Yuwanto and Hanifa Triyuwanti for their patient and support
during my study in Malaysia.
ii
Abstract of thesis presented to the Senate of Universiti Putra Malaysia in fulfilment of the requirement for the degree of Doctor of Philosophy
POPULATION ECOLOGY, REPRODUCTIVE BEHAVIOUR AND FEEDING HABIT OF Helopeltis antonii Signoret ON CASHEW (Anacardium
occidentale L.) PLANTS
By
SISWANTO
November 2007
Chairman : Associate Professor Rita Muhamad, PhD
Faculty : Agriculture
Helopeltis antonii Signoret (Hemiptera: Miridae) is well known as one of the
important pest of the cashew plant, Anacardium occidentale L. Both the
nymph and adult stages feed on young and succulent parts of the plant such
as the young leaves, shoots, inflorescences and fruits causing death of those
parts. This research was conducted with the objectives to establish age-
specific life table, to investigate some biological aspects of H. antonii, to
study the effect of damage caused by H. antonii on shoot, inflorescence and
fruit, and also to study some ecological aspects related to H. antonii
population in the field. The studies were conducted in pesticide-free cashew
plantation belonging to smallholders and also in the laboratory of Estate
Service of Wonogiri Regency in Ngadirojo district, Wonogiri, Central Java,
Indonesia from March 2004 to May 2006. The life table of H. antonii revealed
a high hatchability but a bulk mortality occurred at early nymphal stages and
relatively fewer death during the adult stage. The contribution of the female
towards female births (mx) was at its maximum on the 16th day of oviposition.
iii
The population parameters of H. antonii fed cashew showed that the intrinsic
rate of natural increase (rm) was 0.092/female/day, the net reproductive rate
(Ro) was 12.84, the capacity for increase (rc) was 0.090, the finate rate for
increase ( λ ) was 1.097 female/day, and mean generation time (T) was
27.70 days with the population doubling (DT) every 7.52 days. Biological
studies revealed that H. antonii feeding lesions developed faster on
inflorescence and shoot compared to fruits. The lesions on shoot and
inflorescence produced depressed and wrinkled surface which then dried up
within four days. Meanwhile it caused depression on apple and flattened the
surface on nut. Feeding preference with no choice experiment suggested that
H. antonii preferred to feed on shoot and young fruits rather than the
inflorescence and older fruits. Results in the choice experiment suggested
that H. antonii preferred to feed on shoot compared to inflorescence and
fruits. The female preferred to oviposit on inflorescence compared to shoot
and fruit. The premating period for both male and female H. antonii was one
day. Sex ratio of females to males did not influence the number of eggs laid.
However, overcrowded males seemed to influence female longevity. The
frequency of matings did not influence the number of eggs laid and the
hatchability, eventhough, females which mated more than once tended to lay
more eggs. Damage assessment study revealed that the percentage of
shoot, inflorescence and fruit death increased with the number of lesions.
Apart from the number of lesions, position of lesions and stage (age of part
attacked) also affected damage intensity, particularly on inflorescence and
fruits. Small or young fruits were not able to tolerate heavy damage by H.
antonii, whereas older fruits were relatively not affected by the damage. Field
iv
experiments indicated that cashew fruit particularly small and medium sized
were more susceptible to H. antonii feeding lesions compared to
inflorescences. The infestation of H. antonii was linked to the phenology of
the cashew plants. Higher percentage of inflorescence death occurred in the
second phase of flowering season, meanwhile higher percentage of fruit
death occurred in the third flowering season. Studies on the population
fluctuation and dispersion of H. antonii in cashew plantation indicated that the
fluctuation in the population of H. antonii was cyclical with the population
peaking around July when cashew shoots and inflorescences were
abundant. The population began to increase just after the rainfall season
stopped and reached the peak three months later when rainfall was
intermittent low. Number of shoots and inflorescences of cashew plants had
significant influence on the number of H. antonii. The trend of population
abundance was not directly associated with the rainfall, but rainfall influenced
the physiology of the cashew plant to produced flushes/shoots and
inflorescences. Distribution analysis using various indices of dispersion and
regression models indicated an aggregated distribution when the population
was high during flushing-flowering season of cashew plants and a regular or
random distribution when the population was low during post-flowering
season.
v
Abstrak tesis yang dikemukakan kepada Senate Universiti Putra Malaysia sebagai memenuhi keperluan untuk Ijazah Doktor Falsafah
EKOLOGI POPULASI, PERLAKUAN PEMBIAKAN DAN TABIAT PEMAKANAN Helopeltis antonii Signoret KE ATAS POKOK GAJUS
(Anacardium occidentale L.)
Oleh
SISWANTO
2007
Pengerusi : Profesor Madya Rita Muhamad, PhD
Fakulti : Pertanian
Helopeltis antonii Signoret di kenali sebagai salah satu serangga perosak
pada pokok gajus, Anacardium occidentale L. Kedua-dua nimfa dan dewasa
memakan pada bahagian muda dan bahagian sukulen seperti daun muda,
pucuk, infloresens dan buah yang menyebabkan kematian pada bahagian
ini. Penyelidikan ini dijalankan dengan beberapa objektif iaitu membina
jadual hidup umur-spesifik, untuk mengkaji aspek ekologi H. antonii, untuk
mengkaji kesan kerosakan yang disebabkan oleh H. antonii ke atas pucuk,
infloresens dan buah, dan juga mengkaji aspek ekologi berhubung dengan
populasi H. antonii di lapangan. Kajian ini dijalankan di kawasan pemilik kecil
ladang pokok gajus yang bebas dari pestisid dan di makmal Dinas
Perkebunan Wonogiri di daerah Ngadirojo, Wonogiri, Jawa Tengah,
Indonesia dari Mac 2004 sehingga Mei 2006. Jadual hidup H. antonii
menunjukkan kebolehan menetas yang tinggi tetapi kebanyakan mati pada
tahap awal peringkat nimfa dan relatif rendah pada peringkat dewasa.
Sumbangan betina terhadap kelahiran betina (mx) adalah pada kadar
vi
maksimum pada hari ke 16 peneluran. Parameter populasi H. antonii yang
memakan pokok gajus menunjukkan pertambahan semulajadi kadar intrisik
(rm) adalah 0.092/betina/hari, kadar penghasilan bersih (Ro) 12.84, kapasiti
pertambahan adalah 0.090 dan pertambahan kadar finate (λ) adalah 1.097
betina/hari, dan min tempoh generasi (T) adalah 27.70 hari, dan populasi
akan berganda setiap 7.52 hari. Kajian biologi menunjukkan pemakanan
kecederaan berkembang dengan cepat pada infloresens dan pucuk
berbanding buah. Kecederaan pada pucuk dan infloresens menyebabkan
tekanan dan permukaan berkedut dan kering dalam masa 4 hari, manakala
ianya menyebabkan tekanan pada epal dan permukaan rata pada kacang.
Pemilihan pemakanan dengan eksperimen tiada pilihan menunjukkan yang
H. antonii lebih memilih memakan bahagian pucuk dan buah muda
berbanding infloresens dan buah lebih tua. Daripada kajian eksperimen
dengan pilihan menunjukkan H. antonii lebih memilih memakan bahagian
pucuk berbanding inflorescences dan buah. Betina juga lebih memilih untuk
bertelur pada infloresens berbanding pada pucuk dan buah. Tempoh
pramatang bagi betina dan jantan H. antonii adalah 1 hari. Nisbah jantina
betina kepada jantan tidak dipengaruhi bilangan telur yang dikeluarkan.
Walaubagaimanapun, kelimpahan jantan mungkin mempengaruhi
jangkahidup betina. Kekerapan persenyawaan tidak mempengaruhi bilangan
telur yang dihasilkan dan yang menetas, walaubagaimanapun betina yang
disenyawakan lebih cenderung menghasilkan telur. Kajian penilaian
kerosakan menunjukkan peratus kematian pada pucuk, infloresens dan buah
bertambah dengan bilangan kecederaan. Selain daripada bilangan
kecederaan, kedudukan kecederaan dan peringkat (umur bahagian yang
vii
diserang) juga mempengaruhi intensiti kerosakan terutama pada bahagian
infloresens dan buah. Buah kecil dan muda tidak dapat bertoleransi dengan
kerosakan yang banyak oleh H. antonii, manakala buah tua tidak dipengaruhi
oleh kerosakan tersebut. Kajian lapangan menunjukkan buah gajus terutama
size kecil dan ménengah adalah lebih terdedah kepada kecederaan
pemakanan H. antonii berbanding dengan infloresens. Serangan H. antonii
adalah berkait rapat dengan fenologi pokok gajus. Peratus kematian
infloresens tinggi pada musim kedua berbunga, manakala peratus kematian
buah tinggi pada musim ketiga berbunga. Kajian fluktuasi dan taburan
populasi H. antonii berkitar dengan kemuncak populasi pada sekitar Julai
apabila bilangan infloresens dan buah gajus melimpah. Populasi mulai
bertambah selepas musim hujan berhenti dan mencapai kemuncak tiga
bulan kemudian apabila sela hujan rendah. Bilangan pucuk dan infloresens
pokok gajus sangat signifikan mempengaruhi ke atas bilangan H. antonii.
Corak kelimpahan populasi tidak berhubung langsung dengan jumlah hujan,
tetapi jumlah hujan mempengaruhi fisiologi pokok gajus untuk menghasilkan
pucuk dan infloresen. Analisis taburan menggunakan indeks taburan model
regressi menunjukkan taburan aggregasi apabila populasi tinggi pada musim
berbunga lebat pokok gajus dan biasa atau random apabila populasi rendah
pada akhir musim berbunga.
viii
ACKNOWLEDGEMENTS
All praises and thanks are to Allah SWT., the Almighty, for giving me
guidance, strength and patience in finishing my study. The author invokes
Allah’s blessings of peace for the Holy Prophet Mohammad, the messenger
of Allah.
First of all I would like to express my sincerest thanks and appreciation to
Associate Professor Dr. Rita Muhamad, Chairman of my supervisory
committee for her encouragement, familiar support, invaluable advice and
intellectual guidance during my study, preparation of the research proposal,
in the conduct of the research and in the writing up this thesis. Grateful
thanks are also due to my supervisory committee member, Professor Dr.
Dzolkifli Omar and Dr. Elna Karmawati for their constructive comments,
advice and help throughout my study and encouragement during the
completion of this thesis.
My sincere thanks to the Director General of the Indonesian Agency for
Agricultural Research and Development (IAARD), Dr. Ir. Ahmad Suryana,
MS; The Chair of the Committee of Human Resources Development, Dr.
Haryono; Director of Indonesian Center for Estate Crops Research and
Development, Dr. Bambang Prastowo and the former chair, Dr. Hasnam;
Director of Indonesian Medicinal and Aromatic Crops Research Institute,
Bogor, Dr. Muhamad Syakir and the former Director of Indonesian Spices
ix
and Medicinal Crops Research Institute, Bogor, Dr. Molide Rizal for giving
me the opportunity to take up PhD program.
My gratitude also to Dr. Darmono Taniwiryono, Manager of Integrated Pest
Management for Smallholder Estate Crops Project (IPM-SECP), for the
opportunity and financial support to pursue a PhD study in Univeriti Putra
Malaysia. I would also to thank to all IPM Project staff, especially Mrs. Ina
Purwantini, the financial officer, for their help with financial arrangements.
I would like to thank the Director of Wonogiri Estate Service and Staff for
providing laboratory for my research; cashew farmers in Ngadirojo district,
Wonogiri for permission to use their cashew plots for my experiments,
especially to Mr. Supardi in Pondok village, Ngadirojo district . Thanks are
also extended to Mr. Ahyar, Mr. Cucu Sukmana, Mr. Endang Sugandi and
Mr. Eko Tri Wahyono for their assistance in collecting data for the
experiments. Without their assistance I would have difficulties conducting the
experiments.
I also would like to thank to my housemates especially to Antario Dikin and
Taufik Ratule for sharing the daily joy together during my stay in Malaysia.
Finally, my special thanks to my wife, Yuyun Nurohmah and my children,
Arina Yusianti, Naufal Yuwanto and Hanifa Triyuwanti for their patience,
understanding, support and inspiration given to me during the period of my
study in Malaysia.
x
I certify that an Examination Committee met on 16th November 2007 to conduct the final examination of Siswanto on his degree in Doctor of Philosophy thesis entitled “Population ecology, reproductive behaviour and feeding habit of Helopeltis antonii Signoret on cashew (Anacardium occidentale L.) plants” in accordance with Universiti Pertanian Malaysia (Higher Degree) Act 1980 and Universiti Pertanian Malaysia (Higher Degree) Regulations 1981. The Committee recommends that the student be awarded the degree of Doctor of Philosophy. Members of the Examination Committee are as follows: Kamaruzaman Sijam, PhD Associate Professor Faculty of Agriculture Universiti Putra Malaysia (Chairman)
Yusof Ibrahim, PhD Professor Faculty of Agriculture Universiti Putra Malaysia (Internal Examiner) Rohani Ibrahim, PhD Associate Professor Faculty of Agriculture Universiti Putra Malaysia (Internal Examiner) Idris Abd. Ghani, PhD Professor Faculty of Science and Technology Universiti Kebangsaan Malaysia (External Examiner)
______________________________ HASANAH MOHD. GHAZALI, PhD Professor and Deputy Dean School of Graduate Studies Universiti Putra Malaysia
Date:
xi
This thesis was submitted to the Senate of Universiti Putra Malaysia and has been accepted as fulfilment of the requirements for the degree of Doctor of Philosophy. The members of the Supervisory Committee were as follow: Rita Muhamad, PhD Associate Professor Faculty of Agriculture Universiti Putra Malaysia (Chairman) Dzolkifli Omar, PhD Professor Faculty of Agriculture Universiti Putra Malaysia (Member) Elna Karmawati, PhD Senior Research Officer Indonesian Center for Estate Crops Research and Development, Bogor, Indonesia (Member)
__________________________ AINI IDERIS, PhD Professor and Dean School of Graduate Studies Universiti Putra Malaysia
Date: 21 February 2008
xii
DECLARATION
I hereby declare that the thesis is based on my original work except for quotations and citations which have been duly acknowledged. I also declare that it has not been previously or concurrently submitted for any other degree at UPM or other institutions.
____________________________ Siswanto
Date: 17 - 12 - 2007
xiii
TABLE OF CONTENTS
Page
ii DEDICATION iii ABSTRACT vi ABSTRAK ix ACKNOWLEDGEMENTS xi APPROVAL xiii DECLARATION xvii LIST OF TABLES xix LIST OF FIGURES xxi LIST OF PLATES xxii LIST OF APPENDICES xxiv LIST OF ABBREVIATIONS
Chapter 1 GENERAL INTRODUCTION 1.1 1.1 Background 1.1
2.1 2 LITERATURE REVIEW 2.1 Helopeltis antonii Signoret 2.1 2.1.1 Systematic and morphology of H. antonii 2.1 2.1.2 Biology of H. antonii 2.3 2.1.3 Distribution 2.4 2.1.4 Host plant 2.5 2.1.5 Ecology and fluctuation population 2.7 2.1.6 Distribution pattern 2.9 2.1.7 Damage to cashew 2.11 2.1.8 Control measures 2.14 2.1.9 Natural enemies 2.16 2.2 Cashew (Anacardium occidentale, L.) 2.18 2.2.1 Taxonomy and botany 2.18 2.2.2 Characteristic of cashew 2.21 3 LIFE TABLES AND DEMOGRAPHIC PARAMETERS OF
Helopeltis antonii ON CASHEW 3.1
3.1 Introduction 3.1 3.2 Materials and Methods 3.3 3.2.1 Helopeltis antonii 3.3 3.2.2 Life table construction 3.5 3.3 Results 3.8 3.3.1 Age-specific survival life table 3.8 3.3.2 Age-specific fertility table 3.15 Page 3.4 Discussion 3.18
xiv
3.5 Conclusion 3.22 4 BIOLOGICAL ASPECTS OF Helopeltis antonii 4.1 4.1 Introduction 4.1 4.2 Feeding Behaviour of H. antonii 4.2 4.2.1 Introduction 4.2 4.2.2 Materials and methods 4.2 4.2.2.1 Insects 4.2 4.2.2.2 Location of the experiment 4.3 4.2.2.3 Effect of adult feeding on numbers and
characteristic of lesions 4.3
4.2.2.4 Feeding preference studies 4.3 4.2.3 Results 4.6 4.2.3.1 Effect of adult feeding on numbers and
characteristic of lesions 4.6
4.2.3.2 Feeding preference studies 4.8 4.2.4 Discussion 4.13 4.3 Oviposition Behaviour of H. antonii 4.14 4.3.1 Introduction 4.14 4.3.2 Materials and Methods 4.15 4.3.2.1 Choice of oviposition sites 4.15 4.3.3 Results 4.16 4.3.4 Discussion 4.17 4.4 Mating Behaviour of H. antonii 4.19 4.4.1 Introduction 4.19 4.4.2 Materials and Methods 4.20 4.4.2.1 Premating period of male and female H.
antonii 4.20
4.4.2.2 Influence of sex ratio on fecundity and longevity of female H. antonii
4.21
4.4.2.3 Influence of mating frequency to fecundity and eggs hatchability
4.21
4.4.3 Results 4.22 4.4.3.1 Premating period of male and female H.
antonii 4.22
4.4.3.2 Influence of sex ratio on fecundity and longevity of female H. antonii
4.24
4.4.3.3 Influence of mating frequency to fecundity and eggs hatchability
4.25
4.4.4 Discussion 4.26 4.5 Conclusion 4.28 5 DAMAGE ASSESSMENT OF CASHEW BY Helopeltis antonii 5.1 5.1 Introduction 5.1 5.2 Materials and Methods 5.2 5.2.1 Damage assessment of H. antonii on shoot,
inflorescence and fruit 5.2
Page 5.2.2 Feeding lesions due to natural infestation of H.
xv
antonii on inflorescences and fruits 5.4 5.3 Results 5.5 5.3.1 Damage assessment of H. antonii on shoots 5.5 5.3.2 Damage assessment of H. antonii on
inflorescences 5.8
5.3.3 Damage assessment of H. antonii on fruits 5.10 5.3.4 Feeding lesions due to natural infestation of H.
antonii on inflorescences and fruits 5.18
5.4 Discussion 5.33 5.5 Conclusion 5.35 6 POPULATION FLUCTUATION AND DISPERSION PATTERN OF
Helopeltis antonii 6.1
6.1 Introduction 6.1 6.2 Materials and Methods 6.2 6.2.1 Population fluctuation of H. antonii in cashew
plantation 6.2
6.2.2 Dispersion of H. antonii in cashew plantation 6.4 6.3 Results 6.6 6.3.1 Population fluctuation of H. antonii in cashew
plantation 6.6
6.3.2 Dispersion of H. antonii in cashew plantation 6.14 6.4 Discussion 6.18 6.5 Conclusion 6.23 7 GENERAL DISCUSSION AND CONCLUSION 7.1
REFERENCES R.1 APPENDICES A.1 BIODATA OF THE AUTHOR B.1 LIST OF PUBLICATIONS B.3
xvi
LIST OF TABLES
Table Page 2.1 List of host plants to H. antonii 2.6 3.1 Life table of H. antonii fed shoots of cashew (cohort 1, 20
April 2004). 3.10
3.2. Life table of H. antonii fed shoots of cashew (cohort 2, 22
April 2004). 3.11
3.3 Life table of H. antonii fed shoots of cashew (cohort 3, 22
June 2004). 3.12
3.4 Pooled life table of H. antonii on cashew. 3.15 3.5 Life and age-specific fecundity table of H. antonii fed of
cashew. 3.17
3.6 Population and reproductive parameters of H. antonii fed
shoots of cashew. 3.18
4.1 Number and size of feeding lesion caused by female and
male of H. antonii on different parts of cashew plant. 4.7
4.2 Changes in colour and surface of feeding lesion caused by
adult of H. antonii on different parts of cashew plant. 4.9
4.3 Mean number of feeding lesions on different parts of cashew plant
caused by different stages of H.antonii in feeding preference with no-choice experiment.
4.10
4.4 Mean length of feeding lesions on different parts of cashew
plant caused by different stages of H.antonii in feeding preference with no-choice experiment.
4.12
4.5 Mean width of feeding lesions on different parts of cashew plant
caused by different stages of H.antonii in feeding preference with no-choice experiment.
4.12
4.6 Mean number of lesions on parts of cashew plant caused by
different stages of H. antonii in the feeding preference with choice experiment.
4.13
4.7 Mean number of eggs laid by H. antonii found on different
parts of cashew plant. 4.17
4.8 Fecundity of female H. antonii in premating study of the male
and Female. 4.23
xvii
4.9 Influence of sex ratio on fecundity and longevity of females
H. antonii. 4.24
4.10 Influence of mating frequencies to fecundity and eggs
hatchability of H. antonii. 4.26
5.1 Damage on shoots following exposure to different intensities
of H. antonii infestation at week six. 5.6
5.2 Damage on inflorescences following exposure to different
intentions of H. antonii infestation at weeks six. 5.9
5.3 Damage on different sizes of cashew fruit following
exsposure to different intensities of H. antonii infestation 5.11
5.4 Coefficient of correlation (r) between number of H. antonii
lesions and fruit growth parameters in small, Medium, and large fruits
5.13
5.5 The percentages of fruits that died by H. antonii and
naturally during fruit development in the first, second and third flowering season at week 15.
5.33
6.1 Coefficient of correlation ( r ) between H. antonii population
and their environmental factors during two consecutive flushing-flowering seasons 2004-2005 and 2005-2006.
6.9
6.2 Stepwise regression for H. antonii population against
environmental factors during two consecutive cashew plants season 2004-2006 in Wonogiri, Indonesia.
6.10
6.3 Coefficient of correlation ( r ) between cashew yield and their
environmental factors during two consecutive flushing-flowering seasons 2004-2005 and 2005-2006.
6.13
6.4 Distribution statistics and dispersion indices of H. antonii on
cashew plantation in Wonogiri, Indonesia, 2006. 6.15
6.5 Linear regression for Taylor's Power Law and Iwao's
patchiness regression for seasonal dispersion of H.antonii according to plant phenology of cashew plants in Wonogiri, Indonesia 2004-2006.
6.17
xviii
LIST OF FIGURES
Figure Page 3.1 Patterns of survivorship curve (lx) of H. antonii for three (A, B
and C) different cohorts. 3.10
3.2 Survivorship of adult female and male H. antonii on cashew 3.14 3.3 Daily age-specific survival and fecundity of H. antonii
feeding on Cashew 3.16
5.1 The cumulative percentages of dead shoots during shoot
development following exposure to different intensities of H. antonii infestation
5. 6
5.2 Cumulative percentages of dead inflorescences during
inflorescences development following exposure to different intensities of H. antonii infestation
5.9
5.3 Percentages of H. antonii damage lesions distributed on
stalk, apple and cashew fruits with different level of damage in small (A), medium (B) and large (C) fruits.
5.16
5.4 Percentages of dead inflorescence death during
development caused by H. antonii (H) and not by H. antonii (-H) in the first phase of flowering season.
5.19
5.5 Cumulative number of H. antonii per surviving inflorescence
and the percentages of dead inflorescence associated with H. antonii during the first phase of flowering season.
5.19
5.6 Percentage of dead fruits during fruit development caused
by H. antonii (H) and not by H.antonii (-H) in the first phase of flowering season.
5.21
5.7 Cumulative number of H. antonii lesions per surviving fruit
and the percentages of fruits death caused by H. antonii in cashew during the first of flowering season.
5.21
5.8 Comparison of cumulative mean number of H. antonii
lesions per inflorescence for dead inflorescences due to lesions compared to inflorescences that survived from the lesions during the first phase of flowering season.
5.23
5.9 Mean numbers of feeding lesions from dead inflorescences
at 5-7 weeks old, 8-10 weeks old, 11-13 weeks old and 14-19 weeks old during the first phase of flowering season.
5.23
5.10 Percentage of dead inflorescences during development
xix
caused by H. antonii (H) and not by H.antonii (-H) in the second phase of flowering season.
5.25
5.11 Cumulative number of H. antonii lesions per surviving
inflorescence and the percentages of dead inflorescence caused by H. antonii during the second phase of flowering season.
5.25
5.12 Percentage of dead fruits during development caused by H.
antonii (H) and not by H. antonii (-H) in the second phase of flowering season.
5.26
5.13 Cumulative total number of H. antonii lesions per surviving
fruit and the percentage of dead fruit caused by H. antonii during the second phase of flowering season.
5.26
5.14 Comparison of cumulative mean number of H. antonii
lesions per inflorescence for dead inflorescence due to the lesions compared with inflorescences that survived from the lesions during the second phase of flowering season.
5.28
5.15 Mean numbers of feeding lesions from dead inflorescences
at 5-7 weeks old, 8-10 weeks old, 11-13 weeks old and 14-19 weeks old during the second phase of flowering season.
5.28
5.16 Percentage of dead inflorescences during development
caused by H. antonii (H) and not by H. antonii (-H) in the third phase of flowering season
5.29
5.17 Cumulative number of H. antonii lesions per surviving
inflorescence and the percentages of dead inflorescence caused by H. antonii during the third phase of flowering season.
5.29
5.18 Percentages of dead fruits during development caused by H.
antonii (H) and not by H.antonii (-H) in the third phase of flowering season.
5.31
5.19 Cumulative number of H. antonii lesions per surviving fruit
and the percentages of dead fruit caused by H. antonii in the third phase of flowering season.
5.31
5.20 Comparison of cumulative mean number of H. antonii
lesions per inflorescence for dead inflorescences due to the lesions compared with inflorescences that survived from the lesions during the third phase of flowering season.
5.32
5.21 Mean numbers of feeding lesions fromdead fruits at 5-7
weeks old, 8-10 weeks old, 11-13 weeks old and 14-19 weeks old during the third phase of flowering season.
5.32
xx
6.1 Maps of cashew plant sampling-pattern in Wonogiri, Indonesia, 2004-2006.
6.3
6.2 Population abundance of H. antonii on cashews in relation to
climatic factors (rainfall, temperature and relative himidity) and numbers of shoots and inflorescences in Wonogiri between March 2004 to May 2006.
6.7
xxi
LIST OF PLATES
Plate Page 2.1 H. antonii: (a) thread-like breathing tubes of eggs, (b) young
nymphs, (c) mature nymph and (d) adult. 2.2
2.2 Typical of a cashew tree after the first flushing phase. 2.19 2.3 (A) Inflorescence with opened and unopened flowers, (B)
male flower, (C) perfect/hermaphrodite flower, (D) developing fruit, and (E) fruit: 1. stalk, 2. apple, and 3. nut.
2.20
3.1 New emerged adults fed on cucumber in the laboratory. 3.4 4.1 Framed-tile cage on cashew tree used for the caging
experiments. 4.4
4.2 Caging of branch of cashew consisting shoot, inflorescence
and fruit. 4.5
5.1 Damage on the shoots about six weeks following H. antonii
infestation: (A) slightly damaged, (B) moderately damaged, (C) severely damaged, and (D) control.
5.7
5.2 Damage on the inflorescences about three weeks following
H. antonii infestation:(A) slightly damaged, (B) Moderately damaged, (C) severely damaged and (D) control.
5.10
5.3 Different fruit size of cashew with H. antonii lesions: (A) small
fruit, (B) medium fruit, and (C) large fruit. 5.17
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LIST OF APPENDIX TABLES
Table Page APPENDIX 1 1 Analysis of variance for the number of feeding lesions on
different parts of cashew plant caused by different stages of H. antonii in feeding preference with no-choice experiment
A1
2 Analysis of variance for the length of feeding lesions on
different parts of cashew plant caused by different stages of H. antonii in feeding preference with no-choice experiment
A1
3 Analysis of variance for the width of feeding lesions on
differentparts of cashew plant caused by different stages of H. antonii in feeding preference with no-choice experiment
A1
4 Analysis of variance for the number of feeding lesions on
different parts of cashew plant caused by different stages of H. antonii in feeding preference with choice experiment
A2
5 Analysis of variance for the number of eggs laid by H.
antonii found in different parts of cashew plants A2
6 Analysis of variance for the influence of sex ratio on
fecundity of females H. antonii A2
7 Analysis of variance for the influence of sex ratio on
longevity of females H. antonii A3
8 Analysis of variance for the influence of mating frequencies
to fecundity and eggs hatchability of H. antonii A3
APPENDIX 2 1 Analysis of variance for the shoots following exposure to
different intensities of H. antonii infestation at week six A4
2 Analysis of variance for the inflorescences following
exposure to different intensities of H. antonii infestation at week six
A4
3 Analysis of variance for the length of apple on small cashew
fruits following exposure to different intensities of H. antonii infestation
A4
4 Analysis of variance for the breadth of apple on small
cashew fruits following exposure to different intensities of H. antonii infestation
A5
5 Analysis of variance for the length of nut on small cashew
fruits following exposure to different intensities of H. antonii A5
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infestation 6 Analysis of variance for the breadth of nut on small cashew
fruits following exposure to different intensities of H. antonii infestation
A5
7 Analysis of variance for the wet weight of nut on small
cashew fruits following exposure to different intensities of H. antonii infestation
A6
8 Analysis of variance for the dry weight of nut on small
cashew fruits following exposure to different intensities of H. antonii infestation
A6
9 Analysis of variance for the length of apple on medium
cashew fruits following exposure to different intensities of H. antonii infestation
A6
10 Analysis of variance for the breadth of apple on medium
cashew fruits following exposure to different intensities of H. antonii infestation
A7
11 Analysis of variance for the length of nut on medium cashew
fruits following exposure to different intensities of H. antonii infestation
A7
12 Analysis of variance for the breadth of nut on medium
cashew fruits following exposure to different intensities of H. antonii infestation
A7
13 Analysis of variance for the wet weight of nut on medium
cashew fruits following exposure to different intensities of H. antonii infestation
A8
14 Analysis of variance for the dry weight of nut on medium
cashew fruits following exposure to different intensities of H. antonii infestation
A8
15 Analysis of variance for the length of apple on large cashew
fruits following exposure to different intensities of H. antonii infestation
A8
16 Analysis of variance for the breadth of apple on large
cashew fruits following exposure to different intensities of H. antonii infestation
A9
17 Analysis of variance for the length of nut on large cashew
fruits following exposure to different intensities of H. antonii infestation
A9
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